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DOI: 10.1002/jpln.201200287
355
Review Article
Department of Agricultural Chemistry and Environmental Biogeochemistry, Poznan University of Life Sciences, Wojska Polskiego 71F,
60-637 Pozna, Poland
2 K + S Kali GmbH, 34111 Kassel, Germany
Abstract
The supplies of water and nitrogen to a plant during its critical stages of growth are the main factors that define crop yield. A crop experiences irregular water deficits during its life cycle in rainfed agriculture. An effective anti-stress-oriented approach therefore ought to focus on increasing
the units of water productivity. The main objective of this conceptual review is to confirm that
adequate K management can be used as an important tool to alleviate the negative effects of
water deficit on plant growth, yield-component formation, and yield. The French and Schultz
approach of using the water-limited yield (WLY) was modified in this review into a graphical form
and was used to discriminate between yield fractions that depended on the volume of transpired
water from those that were induced by K fertilizer. By using this method, it was possible to
demonstrate the extent of several crop (winter wheat, spring triticale, maize, sugar beet)
responses to the K supply. Yield increases resulting from K application mostly appeared under
conditions of mild water deficit. As described for sugar beet, finding the critical period of crop K
sensitivity is a decisive step in understanding its impact on water-use efficiency. It has been
shown that an insufficient supply of K during crucial stages in the yield formation of cereals
(wheat, spring triticale), maize, and sugar beet coincides with a depressed development in the
yield components. The application of K fertilizer to plants is a simple agronomic practice used to
increase crop tolerance to a temporary water shortage. It may be that the improvement of a
plants access to K during mild water-deficiency stress will increase water uptake by the root
cells, which in turn increases their osmotic potential and thereby allows extension growth. This
growth in turn promotes access to other mineral elements (including nitrogen) and water, which
favor plant growth and yield.
Accepted April 5, 2013
Key words: drought stress / water-limited yield / critical stage / root system
1 Introduction
The yield potential of a particular crop may be defined as the
maximum plant productivity in the presence of an ample supply of water and nutrients (Evans and Fisher, 1999). However, rain-fed crops are subject to intermittent water stress in
temperate climate zones, which decreases the yield potential,
a trait that is primarily controlled by the genome (Debaeke
and Aboudrare, 2004; Rabbinge and Diepen, 2000). Actual
harvested yields result from the interactions between different
growth factors, which are responsible for fulfilling the yield
potential of cultivated varieties. As a rule, actual yields are
much lower than the attainable maximums and are defined
by soil and climatic conditions (Supit et al., 2010). On the
basis of the degree of yield potential realization, Rabbinge
(1993) has divided all growth factors into three main groups
in descending order of importance: (1) defining (light, CO2,
crop plant), (2) limiting (water, nutrients), and (3) reducing
(weeds, diseases, external toxins). Furthermore, Lansigan
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356
Ya
:
ETa
(1)
(2)
where: Ya, the yield of a particular crop; k, biomass-to-transformation ratio; ETa, total water use (ETa = Tp + Es); Tp, plant
transpiration, Es, soil evaporation; HI, harvest index; VPD,
vapor-pressure deficit.
As a rule, it is assumed that any increase in crop yield in a
particular environment is accompanied by a simultaneous
increase in the WUE (Ritche and Basso, 2008). Therefore,
ETa is frequently considered to be a constant part of the
operational index (Liu et al., 2007). However, Zwart and Bastiaanssen (2004) showed that ETa is not a constant value.
Based on an extended collection of empirical data, these
investigators concluded that the coefficient of determination
or r 2 for the relationship between the harvested yield (Ya) and
ETa was low, amounting to 0.35 for wheat and 0.33 for maize.
In addition, our findings also show that the r 2-coefficient
variability adequately reflects differences in both the climatic
conditions and nutrient supply. Therefore, the WUE is a useful index for comparing different crops and regions (Liu et al.,
2007). Nevertheless, its applicability to discriminating between the growth factors that contribute to yield increases
seems to be limited.
Agnus and Herwaarden (2001) have proposed another method for a simple and quick assessment of water productivity at
the field level, which is known as the French and Schultz
approach (FAS). This method considers the problem of
imbalanced water use by crop plants. Its major thesis
assumes that a plants water supply is not the dominant factor
in determining the quantity of the harvested yield, but
accounts for environmental and management factors as well,
which limit water-use productivity. These assumptions are
seemingly contradictory to the multiple-limitation hypothesis,
(MLH), which treats water as the main growth factor (Rubio
et al., 2003). In fact, the FAS concept raises the problem of
the efficient crop use of water, which in turn depends on facwww.plant-soil.com
RES ;
(3)
WLY TE R
RES WR;
(4)
12000
WLY20+WR
yield of grain, Y / kg ha
-1
10000
8000
WLY20
6000
4000
2000
0
0
100
200
300
400
500
600
precipitation + irrigation / mm
standards of transpiration efficiency
WLY20
WLY20+WR
Figure 1: Graphical interpretation of winter wheat yield response to available water and K against a background with two standards of
transpiration efficiency (basic data adapted from Musolf, 2003).
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358
10000
c
IR
RF
D60
D30
IR
RF
bc
bc
D30
IR
RF
D60
D30
IR
RF
D60
D30
8000
yield of grain / kg ha
-1
6000
4000
2000
-2000
K-
D60
K+
K-
1994
K+
1995
-4000
years and treatments
Yield fraction
WLY20
Y-G
Y-L
Figure 2: Structural components of grain yield from spring triticale as affected by water and K supply (base data adapted from Wyrwa, 1997).
WLY = water-limited yield for TE = 20 kg grain ha1, mm1; Y-G = yield gain, Y-L = yield loss fractional yields; IR = irrigated at BBCH 30 and 60
for 3 weeks; D30, D60 = drought imposed at BBCH 30 and 60, for 3 weeks, respectively, RE = rain-fed conditions. Statistical analysis concerns
real yields; columns marked with the same letters are not significantly different.
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Table 1: WUE indices calculated for spring triticale by two different methods1.
Potassium
Water
treatments
Without
potassium
(K)
With
potassium
(K+)
Available water1
/ mm
mean
WUE-ETa
/ kg ha1 mm1
SD
mean
WUE-FAS
/ kg ha1 mm1
SD
mean
SD
IR
428.6
16.2
13.9
0.07
20.2
3.22
RE
338.7
2.3
11.4
0.14
22.8
0.34
D60
269.9
36.8
7.4
0.01
21.4
24.22
D30
284.1
31.3
5.4
0.07
14.1
14.56
IR
428.6
16.2
15.4
0.07
22.3
3.58
RE
338.7
2.3
15.0
0.64
30.3
1.44
D60
269.9
36.8
13.7
0.64
39.2
19.04
D30
284.1
31.3
11.8
0.71
31.7
22.24
360
12000
de
bc
de
bc
bc
NP.
NPK
NP.
NPK
NP.
NPK
NP.
NPK
10000
-1
8000
6000
4000
2000
0
NP.
-2000
NPK
2005
-4000
2006
2007
2005
sandy loam
NP.
NPK
2006
2007
loamy sand
-6000
soil textural class, years and fertilizing treatments
yield fraction
WLY
Y-G
Y-L
Figure 3: Structural components of maize-grain yields as affected by weather for two soil textural classes and potassium treatments (base data
adapted from Szczepaniak et al., 2010). WLY = water-limited yield for TE = 20 kg grain ha1 mm1; Y-G = yield gain; Y-L = yield loss fractional
yields. Statistical analysis concerns real yields; columns marked with the same letters are not significantly different.
3 Potassium as a water-stress-alleviating
agent
3.1 Determining the critical stage in crop-plant
sensitivity to K supply
The physiological roles of K in plants have been extensively
described in recent reviews (Maathuis, 2009; Marschner,
2012). Potassium is the main osmotic solute in plants and it
affects water homeostasis both at the cellular and the wholewww.plant-soil.com
100000
b
b b
a b
yield of beet / kg ha
-1
80000
60000
Y-L
Y-G
WLY
40000
20000
1996
-20000
1997
1998
1996
L1
1997
198
132
66
198
66
132
198
66
132
198
132
66
198
132
66
198
132
66
0
1998
L2
12
80
10
70
60
50
40
4
30
2
20
0
-2
10
25
45
65
85
105
125
145
165
0
-4
362
-10
days from emergence / d
K characteristics:
AR-K
RRA-K
period would extend from the 72nd to the 95th day of sugar
beetvegetation development. The period of highest demand
in the sugar beet canopy for K coincides with its main growth
parameters. Firstly, the absolute rate of dry-matter and nitrogen accumulation in both the leaves and roots by a highyielding sugar beet canopy takes place between 80 and 90
days for sugar beet vegetation (Grzebisz et al., 2012, Szczepaniak et al., 2012). The relative rate of K accumulation
(RRAK) is another crop characteristic, which is useful in
determining the earliest stage of its sensitivity to a particular
nutrient supply. In the present case, the RRAK reached a
maximum around the 7th week after plant emergence. At this
period of sugar beet development, the rate of foliage expansion depends on the supply of nitrogen, which is a decisive
factor in solar-energy capture (Malnou et al., 2006). According to Marschner et al. (1996) the rate of nitrate-ion uptake is
significantly related to a concomitant K supply. Both parameters, i.e., the maxima for the absolute rate of K accumulation (ARK) and the relative rate of K accumulation (RRAK),
define the critical period of a crops response to K supply.
Both K-accumulation parameters are most likely inherently
related to the rate of root-system extension (length, RL) during the growth season. The key characteristics of root-system
growth during the growing season are the transition point
(TPRL) and the rate of extension (RLr). Both parameters have
been calculated for sugar beets (based on Windt, 1995) by
using the first derivative of the polynomial function of the third
degree (RL):
RL = 0.012d 3 + 3.022d 2 76.26d 379.3 for R2 = 0.99,
RL = 0.036d 2 + 6.044d + 76.26,
3.2.1 Wheat
The majority of arable lands worldwide are cropped with cereals (FAOSTAT, 2012). The sensitivity of this group of crops
to K is generally weakly recognized at the canopy level. The
main reason for this knowledge gap is the high genetic adaptability of cereal species to varying soil fertility and water environments (Calderini and Slafer, 1999; Zhang et al., 2009). In
Europe, wheat is among the most productive cereals, but it
also requires the most fertile soils (Rabbinge and Diepen,
2000). The yield potential of this crop in Europe is high, ranging from 7.7 t ha1 in Bulgaria to 12.4 t ha1 for Ireland, in
turn indicating a slight decrease following the increasing
impact of the continental climate. However, actual yields
show a much deeper contrast between European countries
(Olesen and Bindi, 2002; Supit et al, 2010). The size of the
yield gap depends upon both the growth conditions (soil and
climate) and nutrient management, especially K (Grzebisz
and Fotyma, 2007; Grzebisz and Diatta, 2012; Olesen and
Bindi, 2002; Rabbinge and van Diepen, 2000).
A winter wheat yield of 10 t ha1 exerts the highest demand
for K during the period that extends from the beginning of
shooting (BBCH 30) to the beginning of anthesis (BBCH 60;
Cannell, 1984). This period lasts approximately 45 d, and the
daily rate of K uptake amounted to 4.4 kg K ha1. In this case,
the maximum rate of K accumulation occurred at heading,
and then slightly decreased towards anthesis. At this particular stage of growth, wheat reached maximum K accumulation
of 320 kg K ha1. In addition, its K uptake always preceded
the amount of simultaneously accumulated nitrogen. Therefore, the pattern of K accumulation in wheat during a given
season can be considered to be a prerequisite of high grain
yield. This conclusion is supported by a wheat study from
Great Britain. As reported by Barraclough (1986), the daily
rate of nitrogen and K uptake of 2 and 3 kg ha1 during the critical period (180260 d after sowing) is required to cover
requirements for the production of 9.5 t ha1 of wheat grain.
As reported by King et al. (2003), the shoot-dry-mass
increase is about sevenfold during this period, which indirectly underlines a high sensitivity to the shortage of both
nutrients. However, the impact of water shortage on wheat
performance in the British Islands is much lower than in Central Europe or in semiarid regions (Dodd et al., 2011; Rabbinge and Diepen, 2000; Turner, 2004). The main reason for
this yield gap is the frequency and severity of drought during
wheat development (Passioura, 2006).
Figure 6 shows the results of simultaneously imposed water
and K stress, which significantly affected the K-accumulation
rate (ARK) of wheat during the growth season. For most treatments, the highest ARK was achieved by wheat at the booting
stage (BBCH 40-49). With an ample water supply, the
recorded growth rate of wheat plants at this stage did not
depend on the soil content of available K. Therefore, it can be
concluded that an adequate water supply during the most critical stages of wheat growth (from shooting to anthesis) can,
to some extent, be alleviated by additional water. In many
parts of the world, irrigation is the key to ameliorating waterdeficiency stress, and it significantly increases wheat yields,
i.e., WUE (Liu et al., 2007). As shown in Fig. 6, plants experiencing a water shortage developed different K-accumulation
strategies. Those grown on K-rich soil could maintain a sufficiently high rate of K accumulation during most of the season.
However, plants experiencing both water and K stresses had
a decreasing trend in their K-accumulation rates from the
booting stage onwards.
In light of the known K-accumulation patterns for wheat, a
question arises about the response of yield structure compo-
-1
-1
7
6
LSD0.05
5
4
3
2
IR/D
IR/D
BBCH 30-49
BBCH 61-80
0
20
40
treatments:
61
82
days from spring regrowth / d
K- IR
K- D
K+ IR
103
141
K+ D
Figure 6: Potassium-accumulation rate in wheat under different water and K-fertilizer treatments (basic data adapted from Musolf, 2003). KIR,
K+IR = irrigated treatments without and with K application; KD, K+D = treatments with double drought imposed for 3 weeks during stem
elongation and anthesis, without and with K application.
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364
R2
= 0.51, n = 9,
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ha1, was achieved in 1998; however, the yields did not show
any response to increasing K rates. Analogous conclusions
have been drawn by Milford et al. (2000) under British conditions. The same trend of seasonal beet response to applied K
was reported by Damm et al. (2013).
The study on dry-matter accumulation by sugar beets in response to K management showed a high dependence on the
weather pattern. As presented in Fig. 8, the absolute rate of
beet growth was affected both by the weather within a given
year and K fertilization. In 2001, the water supply during the
summer months amounted to 235 mm, while it was only
105 mm in 2003. The absolute rate of dry matter for beet
growth was higher in the NPK treatment, regardless of the
weather. The degree of difference between non-K and K-fertilized treatments was much higher in 2001, when the water
supply was ample. However, during the dry conditions of
2003, the maximum rate of beet and absolute growth was significantly delayed. This finding is most likely the key reason
for yield depression in relation to the summer water deficit.
A study on the response of sugar beets to induced water and
K stress during the summer months showed a significant dependence on the K supply. As reported by Musolf et al.
(2004b), plants grown under conditions without K application
had much lower concentration of K and nitrogen, both in the
leaves and young roots, at the 6th-leaf stage. It is worth mentioning that an insufficient supply of nitrogen in beets results
in a slower rate of canopy growth. As a consequence, plants
are not capable of efficiently absorbing solar energy, which is
critical to their final yield (Malnou et al., 2006; Milford et al.,
1985). Early K stress impacts the performance of the sugar
beet canopy, which is the primary reason for its high sensitivity to water stress. As presented in Tab. 2, the artificially
imposed water stress affected K accumulation in plants in
-1
-1
50
45
40
35
30
25
20
15
10
5
0
40
57
77
92
113
days from sowing / d
NPK2001
NP2001
134
NPK2003
155
175
NP2003
Figure 8: Effect of natural drought and limiting K supply on absolute rate of storage-root growth in sugar beet (adapted from Grzebisz et al.,
2012).
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366
Table 2: Accumulation of K by sugar beet around the transition point (TP) of canopy growth as imposed by different water and potassium
regimes in kg ha1.
Water treatments Leaves
Storage roots
Total
K+
K+
K+
Rain-fed
216.9ab
88.9c
61.7a
20.1d
278.7ab
109.1c
Irrigation
237.0a
78.4c
55.0b
16.5d
292.1a
94.9c
Drought
172.5b
88.0c
43.5c
18.9d
219.1b
106.9c
based on Musolf (2003); numbers marked with the same letters are not significantly different. Irrigation and drought were imposed in July.
4 Potassium as a root-growth-induction
factor
In general, the pattern of root-system development during the
growth season for particular crop species is consistent and
under genetic control (Taylor and Klepper, 1978). However,
this pattern undergoes changes throughout consecutive
vegetative stages that depend on the soil physical properties,
water and mineral-nutrient content throughout the soil profile
(Smucker and Aiken, 1992). Soil-profile moisture plays a dual
role and is considered to be a plant-growth resource and the
source of life for growing roots. The rate of nutrient uptake by
crop plants is governed by the rate of growth, so the demand
is stage-dependent. In general, it is assumed that roots grow
intensively in the top soil at the beginning of the growth period. As a rule, this soil layer contains sufficient amounts of
nutrients to cover the requirements of the developing seedlings (King et al., 2003). In contrast, the amounts of water
and available nutrients in the upper soil layer continuously
decreases during crop development, which forces plants to
extend their roots towards moist and nutrient-rich zones of
the soil profile (Forde and Lorenzo, 2001). Therefore, the
2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
23,4
23,2
Y18-24 = 0.013K + 19.67
R2 = 0.83, P < 0.01
23
22,8
22,6
22,4
22,2
22
21,8
21,6
21,4
21,2
100
120
140
160
180
200
220
240
260
0-6
18-24
Figure 9: Available water capacity as a function of soil available K content (basic data adapted from Damm et al., 2013).
Crops that are cultivated under rain-fed conditions are subject to frequent wettingdrying cycles during the vegetative
period. Under these conditions, nutrient and water supply to
roots on one side and their growth rate on the other side are
significantly disturbed. The main issue of interest is the
impact of a particular nutrient on root extension. Most published scientific papers on this subject focus on nitrogen and
phosphorus (King et al., 2003; Van Noordwijk and Geijn,
1996). There are limited data on K, especially regarding its
impact on the root system when the plant is subjected to a
water deficit. In addition, reports dealing with the K supply to
a crop plant and the resulting root-system responses are contradictory. The most frequently cited work by Tennant (1976)
implicitly indicated that a lack of K in the growth medium of
wheat significantly decreased the number of seminal roots
and their length, and at the same time entirely halted the development of nodal roots. In addition, the root growth, especially in nodal roots, were considerably reduced in response
to a half rate or a doubled rate of the required amount of K.
These conclusions are contradictory to results presented by
Drew (1975), who showed prolific growth of barley roots in response to increasing rates of applied K.
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368
0,1
depth of rooting / cm
0,2
0,3
0,4
0,5
0,6
0,7
Y/K+ = 45.084e
R2 = 0.96
10
20
KK+
30
Y/K- = 42.489e
R2 = 0.94
40
50
60
URRW IUDFWLRQ
18
b
K/N = 1.8
16
accumulation of nutrient / kg ha
-1
14
12
10
nutrients
K/N = 1.5
8
6
4
2
0
K-
K+
potassium treatments
Figure 11: Effect of long-term K fertilization on nitrogen and K accumulation in sugar beet at BBCH 16 (basic data adapted from Musolf, 2004b).
Analysis of variance conducted for each nutrient separately; respective columns marked with distinct letters are significantly different.
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growth were revealed during the year with prevailing semidrought conditions. As shown in this paper, all crops cultivated in plots with high content of both available and recently
applied K grew faster than the control K treatments, i.e., with
medium attainable K content and without freshly applied K.
As shown in Fig. 10, the fractional distribution of sugar beet
true roots at the full-rosette stage showed a significant response to differences in soil K status. Plants grown on soil
that was rich in available K could increase the fraction of
actual roots in the deeper soil layers as a consequence of the
faster growth rate. This phenomenon is a classic example of
a drought-avoidance strategy that was used by sugar beet
plants in response to a mild water-stress deficiency (Farooq
et al., 2009; Monneveux and Belhassen, 1996; Shao et al.,
2009).
5 Concluding remarks
Water-deficiency stress is irregular in temperate regions,
which has unforeseeable results throughout crop-plant development of varying duration and severity (Ludlow and
Muchow, 1990; Monneveux and Belhassen, 1996). Therefore, the efficient water management of any farming system
should be oriented towards increasing the resistance of crops
to potential water stress. However, the degree of a crop
plants response to water deficit is stage-dependent (Martyniuk, 2008). Yield losses occur when water stress negatively
impacts the dry-matter-accumulation rate, in turn inducing a
change in its partitioning among the plant organs during development progress (Bingham, 2001; Farooq et al., 2009). As
a consequence of water deficits, the development of key yield
components is disturbed, so it has been believed that any
growth factor that increases the uptake of water and/or its
internal efficiency use can decrease the extent of waterstress effects on the plant-growth rate (Debaeke and Aboudrare, 2004; Ritche and Basso, 2008). Potassium is a plantgrowth factor that should be carefully considered on the basis
of its physiological function in the plant, and as one of the
most important amendments that can be applied by farmers
to cope with water-stress deficiency (Cakmak, 2005).
This review has evaluated the alleviating effects of K application on the basis of yield increases. The positive response of
selected crops to K fertilizer is mostly revealed under conditions of mild water deficit. The drought resistance increase is
2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
370
Acknowledgments
A decreased soil water content interferes with relationships
between cations in the soil solution, which in turn modifies
the intracellular concentration of Ca2+ ions in the root apoplast (Shao et al., 2007). Calcium acts as a sensor of environmental signals, including the soil-water gradient, which
affects the processes that are responsible for plant water
management (Bray, 1997; Luan, 2002). Potassium deficiency
in the growth medium also affects the Ca2+ sensors, which in
turn activate the high-affinity K+ transporters in cells within
the root cortex (Maathuis, 2009; Szczerba et al., 2009).
Recent investigations have also revealed that nitrate and K
deficiency can be recognized in the growth medium by the
same molecule, protein kinase (CIPK23). Its activation by calcium-binding proteins is a required step to induce the highaffinity K transporters (Ho and Tsay, 2010).
The nitrogen and especially the nitrate status of a plant is the
key factor in regulating a series of processes that are responsible for the synthesis, signaling, and control of hormone
flows in the shootroot continuum. The response of young
parts of the plant to an insufficient supply of nitrogen results
in a decreased downward transportation of auxin (Krouk
et al., 2011). The movement of this signal molecule within the
roots has been addressed by K carrier TRH1 (Vicente-Agullo
et al., 2004). As a consequence, this molecule affects the
rate of growth and the shape of the root system in the soil
profile (Ashley et al., 2006; Casson et al., 2003; Eapen et al.,
2005).
The modified root system of a crop plant is a consequence of
a consecutive series of metabolic and physiological processes, which result from the action of hormones in the root
shoot continuum. Therefore, these secondary processes,
which are primarily induced by nutrient deficiency in the plant
medium, have recently been termed as trophomorphogenic
plant response (Forde and Lorenzo, 2001). There is most
likely some cross-talk between water- and nutritional-deficiency signaling. The water supply to a plant is considered to
be the main factor in nutrient mass flow, and can induce plant
nutrient-deficiency signaling, which in turn is a driving force
for root growth into the soil layers to reach both water and
nutrients.
The proposed hypothesis of water-stress deficiency control
relies on a simple agronomic trait, i.e., the improvement of
soil-K management. It is well recognized that a decrease in
soil water content is accompanied by a subsequent decrease
in transpiration rate, resulting in less mass flow of K to the
root surface, which in turn reduces K uptake in the K-nonamended soil. Increasing K in the soil solution through fertilizer
application compensates for this effect by providing an adequate K supply and maintaining root growth. Maintaining root
growth allows access to other mineral elements (including
nitrogen) and water. Thus, yields are greater under mild
drought when K fertilizer is added. High available K content in
2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
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